Pristimantis daquilemai, Brito-Zapata & Reyes-Puig & Cisneros-Heredia & Zumel & Ron, 2021
Brito-Zapata, David, Reyes-Puig, Carolina, Cisneros-Heredia, Diego, Zumel, Daniel & Ron, Santiago R., 2021, Description of a new minute frog of the genus Pristimantis (Anura: Strabomantidae) from Cordillera del Condor, Ecuador, Zootaxa 5072 (4), pp. 351-372: 355-365
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Pristimantis daquilemai sp. nov.
Common names. English: Daquilema’s Rain Frog. Spanish: Cutín de Daquilema.
Holotype. ZSFQ 1206, adult female collected near the community of Río Blanco (3.907196°S, 78.485674°W, 2054 m), Paquisha parish, Paquisha county, province of Zamora Chinchipe, Republic of Ecuador ( Fig. 4 View FIGURE 4 ), collected on 10 February 2018 by David Brito-Zapata and Juan Hurtado. GoogleMaps
Paratypes (22 ♀, 22 ♂). Adult females collected at the type locality by David Brito-Zapata and Juan Hurtado : ZSFQ 1201 (3.906969°S, 78.485297°W, 2044 m) GoogleMaps ; ZSFQ 1202–1203, ZSFQ 1207 (3.907441°S, 78.485835°W, 2041 m) GoogleMaps ; ZSFQ 1204–1205, ZSFQ 1210 (3.906679°S, 78.484964°W, 2059 m) GoogleMaps ; ZSFQ 1208 (3.906851°S, 78.485045°W, 2050 m) GoogleMaps , on 10 February , 2018 ; DHMECN 13055 (3.906607°S, 78.484857°W, 2073 m) GoogleMaps , on 12 January, 2016.Adult females collected at La Herradura (4.054202°S, 78.556270°W, 1750 m), Chinapintza , Paquisha county, province of Zamora Chinchipe, Ecuador, by Manuel A. Morales on 30 June, 2003 GoogleMaps : QCAZ 30842–30843 View Materials , QCAZ 30847– 30848 View Materials , QCAZ 30852 View Materials , QCAZ 30857–30859 View Materials , QCAZ 30861–30863 View Materials , QCAZ 30868–30869 View Materials . Adult males collected at the type locality by David Brito-Zapata y Juan Hurtado : ZSFQ 1209 (3.907441°S, 78.485835°W, 2041 m) GoogleMaps ; ZSFQ 1211 (3.906516°S, 78.484839°W, 2054 m) GoogleMaps ; ZSFQ 1212 (3.906589°S, 78.484893°W, 2067 m) GoogleMaps , on 10 February , 2018 ; DHMECN 13066 (3.910324°S, 78.500900°W, 1740 m) GoogleMaps , DHMECN 13071, (3.892927°S, 78.516816°W, 1554 m) GoogleMaps , DHMECN 13080–13081, 13059 (3.906607°S, 78.484857°W, 2073 m) GoogleMaps , on 12 January , 2016. Adult males collected at La Herradura, by Manuel A. Morales on 30 June, 2003 : QCAZ 30844–30846 View Materials , QCAZ 30849–30851 View Materials , QCAZ30853–30856 View Materials , QCAZ 30860 View Materials ; QCAZ 30865–30867 View Materials .
Etymology. The specific epithet “daquilemai” is a noun in the genitive case and a patronymic for Fernando Daquilema Guamán, an indigenous of the Puruhá culture from Ecuador, who led an insurrection in December 1871, considered as one of the most important indigenous rebellions in 19 th century, Ecuador. Known as the Yaruquíes insurrection, it was caused by high church taxes and forced labor for national roadworks. The movement mobilized thousands of people against the tactics of the Ecuadorian state-making model carried forward by the presidency of Gabriel García Moreno ( Ibarra, 2018; López-Ocón, 1986; Mutlak, 2009).
Diagnosis. Pristimantis daquilemai differs from its congeners by the following combination of characters: (1) Dorsal skin shagreen with scattered small tubercles, flanks densely tuberculated; “› ‹”-shaped scapular folds, with two subconical tubercles on the medial and posterior sections of folds; with thin dorsolateral folds extending from scapular folds to post-sacral region; venter areolate; discoidal fold weakly defined; (2) tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally, hidden by muscle and skin; supratympanic region with a weakly defined fold, conspicuous conical or subconical postrictal tubercles; (3) snout subacuminate in dorsal view, rounded in profile, with a prominent rostral papilla at tip of snout; (4) upper eyelid with one elongated conical tubercle and several conical and subconical smaller tubercles, inter-ocular distance wider than upper eyelid; cranial crests absent; (5) dentigerous processes of vomers present, oblique, with two or three teeth; (6) males with weakly defined nuptial pads on dorsum of Finger I; vocal slits and vocal sac absent; (7) Finger I shorter than Finger II; emarginated discs of fingers broadly expanded on fingers II–IV; (8) fingers with lateral fringes; (9) ulnar tubercles present, conspicuous, conical; (10) heels with a conical tubercle; two or three outer tarsal subconical tubercles; indistinct inner tarsal fold; (11) inner metatarsal tubercle ovoid, about 3–4x the size of subconical, rounded, outer metatarsal tubercle; (12) toes with lateral fringes; supernumerary plantar tubercles present, weakly defined basal membrane; expanded discs on all toes, expanded in the same proportion as discs of fingers; Toe V longer than Toe III (disc of Toe V not reaching distal subarticular tubercle on Toe IV; (13) in life, dorsal background orange-brown to dark brown with numerous little orange spots with brown markings; venter and hidden surfaces of posterior legs from greenish-yellow to dark brown with small white dots; groin with orange or yellow spots, with highest intensity and size in females; iris golden with black reticulations and a horizontal reddish brown bar (14) SVL in females 17.3± 1.1 mm (15.5–19.0 mm, n = 22), in males 13.2± 0.9 mm (12.0–15.0 mm, n = 22).
Comparisons. Detailed comparisons with similar species are presented in Table 1 View TABLE 1 . Pristimantis daquilemai is morphologically similar to P. trachyblepharis ( Boulenger, 1918) , P. aquilonaris (Edgar Lehr, Aguilar, Siu-Ting, & Carlos Jordán, 2007), P. minimus Terán-Valdez & Guayasamin, 2009 , P. altamnis Elmer & Cannatella, 2008 , and P. kichwarum Elmer & Cannatella, 2008 . P. daquilemai and P. aquilonaris have their upper eyelids with a conical tubercle and some smaller tubercles and colored pattern in inner surfaces of legs. Males of P. daquilemai , P. trachyblepharis and P. aquilonaris lacking vocal slits. All four species are easily differentiated from P. daquilemai by having a visible tympanum, which is not visible in P. daquilemai . Pristimantis minimus resembles the new species by its small size and by lacking an external visible tympanum. However, both species can be distinguished by the presence of a prominent rostral papilla at the tip of the snout in P. daquilemai (absent in P. minimus ). Pristimantis aquilonaris (females SVL = 19.4–2.0 mm and males SVL = 13.7–17.6 mm), P. kichwarum (females SVL = 24.4–27.0 mm and males SVL = 17.5–21.9 mm) and P. altamnis (females SVL = 26.9–27.6 mm and females SVL = 16.9–19.9 mm) are larger than P. daquilemai (females SVL = 15.5–19 mm and males SVL = 12.0–15.0 mm). Pristimantis aquilonaris , P. kichwarum and P. altamnis have W scapular folds, while P. daquilemai has “› ‹” scapular folds. Pristimantis trachyblepharis present low ulnar tubercles and P. altamnis lack ulnar tubercles (conical and conspicuous in P. daquilemai ). In P. trachyblepharis the rostral papilla is much less evident, no conspicuous colors on groin or other hidden surfaces, contrary to the new species. P. daquilemai has groin with orange or yellow spots, which may extend to the inner surfaces of legs and flanks, while P. aquilionaris has groin, anterior and posterior surfaces of thighs, concealed surfaces of tarsus, and axilla blackish brown with yellowish-orange or reddish-orange flecks ( P. trachyblepharis , P. kichwarum and P. altamnis without colored pattern in inner surfaces).
Other species closely related are in a clade including P. parvillus ( Lynch, 1976) , P. nietoi Arteaga, Pyron, Peñafiel, Romero, Culebras, Bustamante, Yánez-Muñoz and Guayasamin, 2016 , P. walkeri ( Lynch, 1974) , P. luteolatralis ( Lynch, 1976) and P. ceuthospilus ( Duellman & Wild, 1993) . Pristimantis daquilemai and P. ceuthospilus have blotches on groins and inner surfaces of legs, P. ceuthospilus lacks scapular folds, evident rostral papilla and subacuminate snout (present in P. daquilemai ), also bears evident tympanic annulus and membrane (completely covered by skin in P. daquilemai ), and has larger SVL: females SVL = 25.1 mm, 23.5–26.7 mm, and males SVL = 22.4 mm, 19.0– 25.8 mm ( Pristimantis daquilemai : females SVL = 17.3± 1.10 mm, 15.5–19 mm, males SVL = 13.19± 0.87 mm, 12.0–15.0 mm). For the remaining species ( P. parvillus , P. nietoi , P. luteolateralis and P. walkeri ) the main characters that allow to differentiate P. daquilemai from these species are tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally and presence of an elongated conical tubercle on the upper eyelid (tympanic annulus evident and lack of elongated conical tubercle on the upper eyelid in the aforementioned species). Another species that could be similar to P. daquilemai due to its small body size is P. andignomus Lehr & Coloma, 2008 , however, the latter does not present orange-yellow spots in the groin, unlike P. daquilemai .
Description of holotype. Adult female (16.8 mm SVL, Fig. 2–3 View FIGURE 2 View FIGURE 3 ), head as long as wide (40.6% of SVL); snout long (EN 15.6 % of SVL, eye-nostril distance equal to eye diameter), subacuminate in dorsal view, rounded with a prominent rostral papilla on the tip in profile ( Fig. 3 View FIGURE 3 ); nostrils slightly protuberant, directed dorsolaterally ( Fig. 3 View FIGURE 3 ); canthus rostralis weakly concave in dorsal and lateral view; loreal area slightly concave, lips rounded; upper eyelid with one enlarged conical tubercle surrounded by several smaller, conical and subconical tubercles ( Fig. 3 View FIGURE 3 ); interorbital area flat (EW 80% IOD); cranial crests absent; tympanic membrane not differentiated from surrounding skin, tympanic annulus not visible externally, hidden by muscle and skin; supratympanic fold weak, conspicuous conical or subconical postrictal tubercles. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomer small, ovoid, separated, posteromedial to choanae, with 3 teeth. Tongue longer than wide, notched posteriorly, posterior 30% not adhered to mouth floor.
Dorsal skin shagreen with scattered small tubercles, “› ‹”-shaped scapular fold present, with two subconical tubercles on medial and posterior sections of folds, thin dorsolateral folds extending from scapular folds to the post-sacral region; flanks densely tuberculated; skin on chest and venter areolate; discoidal fold weakly defined; cloacal region with two large rounded tubercles. Two conical outer ulnar tubercles; outer palmar tubercle bifid, approximately two times the size of elongate inner palmar tubercle; supernumerary tubercles present; subarticular tubercles well defined, rounded in lateral view. Fingers with narrow lateral fringes; Finger I shorter than Finger II; emarginated discs of fingers broadly expanded, most prominent on Fingers II–IV, about 2× the size of the digit; discs rounded terminally, all with ventral pads well defined by circumferential grooves ( Fig. 5 View FIGURE 5 ).
Hind limbs slender (TL 56.3% of SVL; FL 43.7% of SVL); heel with one conical tubercle, inner edge of tarsus with two or three subconical tubercles; inner metatarsal fold indistinct; toes with narrow lateral fringes, with basal membrane, distinctive between toes IV and V; subarticular tubercles round; inner metatarsal tubercle ovoid, about four times the size of subconical outer metatarsal tubercle; plantar supernumerary tubercles present ( Fig. 5 View FIGURE 5 ); discs of toes expanded in all discs, slightly narrower than those on fingers; toes with ventral pads well-defined by circumferential grooves; Toe V longer than Toe III, disc of Toe V not reaching the base of distal subarticular tubercle on Toe IV ( Fig. 5 View FIGURE 5 ).
Coloration of holotype in preservative. Dorsum grayish-cream with irregular brown blotches, head with post-occipital W- shaped cream mark, “› ‹”-shaped scapulars black marks; flanks and dorsal surfaces of the limbs light brown with transversal bars dark brown; dorsal surfaces of hands and feet thickly dotted with grayish-brown; throat and venter cream with scattered irregular dark brown blotches, aggregated in the throat; little white spots concentered in the venter; groins and inner surfaces of thighs with pale pink blotches delimited by dark brown, extending towards the flanks; ventral surfaces of limbs cream with dark brown blotches; outer edge of the tarsus with a dark brown longitudinal stripe; dark brown cloacal mark ( Fig. 2 View FIGURE 2 , 7 View FIGURE 7 ).
Coloration of holotype in life. Dorsum light brown with irregular dark brown blotches, dorsolateral folds orange, head with “› ‹”-shaped black scapular folds delimited by orange; flanks with brown bars separated by greenish-yellow, scattered tubercles pigmented with orange; venter light brown with minute dark brown spots and white small low warts, throat light brown with scattered irregular dark brown blotches; ventral surfaces of limbs light brown with minute dark brown and orange spots, ventral surfaces of hands and feet light brown, supernumerary and subarticular tubercles pigmented with orange; groin and inner surfaces of thighs and calves with distinctive orange blotches; snout light brown with weakly defined light brown subocular bars; golden iris with black reticulations and a horizontal reddish brown bar; ()-shaped dark brown cloacal mark ( Fig. 6 View FIGURE 6 ).
Measurements (in mm) of holotype: SVL= 16; E–N= 2.5; HL= 6.5; HW= 6.5; IOD= 2.5; IND= 2.5; TL= 9; FL= 7; HL= 45; ED= 2.5; EW= 2.0.
Variations. Measurement ranges and proportions are showed in Table 2 View TABLE 2 . Snout-vent length in females from 15.5 to 19.0 mm and males from 12.0 to 15.0 mm. Tubercles on the scapular fold and dorsolateral folds visible in life and reduced in preservative. Dorsum varies from cream (e.g., ZSFQ 1211), grayish cream (e.g., ZSFQ 1201), grayish-brown (e.g., ZSFQ 1206), gray (e.g., ZSFQ 1204), light brown (e.g., QCAZ 30858) to dark brown (e.g., ZSFQ 1212), with irregular dark brown marks. Males with black “› ‹”-shaped scapular folds except one (DHMECN 13071) in which they are depigmented. In addition, male QCAZ 30855 has a dorsal dark brown stripe, and the male DHMECN 13059 has a dark brown facial mask extending from the nostril to the scapular region. Males with weakly defined white nuptial pads on dorsum of Finger I.
Females are more variable; some of them with a dorsal grayish-brown stripe delimited by dark brown (e.g., DHMECN 13055), others with “˄”-shaped grayish-brown mark in the sacral region (e.g., ZSFQ 1201). Female QCAZ 30852 has the same facial mask as male DHMECN 13059. Meanwhile, the female QCAZ 30863 has a longitudinal brown stripe on dorsum and a gray helmet-shaped mark in the post occipital region. All females have black scapular folds except QCAZ 30858 in which they are semi pigmented. Ventral coloration varies from cream (e.g., ZSFQ 1211), grayish cream (e.g., ZSFQ 1204) to light brown (e.g., QCAZ 30852). Ventral pattern varies from dotted with dark brown (e.g., DHMECN 13071), to densely mottled (e.g ZSFQ 1204, QCAZ 30852). In preservative, all females have groins with pale pink marks delimited by dark brown. These marks are inconspicuous in males ( Fig. 7 View FIGURE 7 ; Fig. 8 View FIGURE 8 ).
Distribution and natural history. This species is known from three localities at elevations between 1554 and 2067 m, on the Cordillera del Condor, southern Ecuador. All localities are geopolitically in the Paquisha county, province of Zamora-Chinchipe. The ecosystem at type locality belongs to Montane Evergreen Forest of the Cordilleras del Cóndor and Kutukú (BsMa02) (MAE, 2013). It is considered a peculiar dwarf forest having plant species of foothills and lower montane forest of the same mountain range but smaller in size. This ecosystem shows a high abundance of epiphytes, bryophytes, and leaflitter, understory is dominated by Chusquea and canopy reach up to 12 m ( Morales et al. 2013). Pristimantis daquilemai was found active at night on leaflitter and in low vegetation, up to 0.6 m above ground. Stomach contents of specimen ZSFQ 1203 consisted of one ant, subfamily Dolichoderinae (Hymenoptera) , one wasp ( Hymenoptera ), and remnants of one fly (Diptera). Syntopic anurans included P. quaquaversus , P. ledzeppelin , and four unidentified species of Pristimantis .
We follow IUCN criteria (Gärdenfors, Hilton-Taylor, Mace, & Rodríguez, 2001). All known records of Pristimantis daquilemai are from the Cordillera del Condor. Habitat change and loss across this mountain range are increasing due to mining activity and two of the three known localities are within mining concessions. These concessions are already registered and are in an active process of exploration and, based on their practices, correspond to largescale mining concessions (IGF 2019). We make the following extinction risk assessment for P. daquilemai : (1) We suspect that P. daquilemai will suffer population declines due to continuous decline in habitat quality across its distribution, however, we refrain from use criterion A until more ecological data are available. (2) P. daquilemai has an estimated extent of occurrence EOO of 35.3 km 2, within the threshold for Critically Endangered (<100 km 2) under criterion B1. If the species occurs in additional sites but is still restricted to the Cordillera del Condor, its eoo would be <2500 km 2, within the threshold for Endangered (<5000 km 2). (3) The species has an estimated AOO of 12 km 2, which is within the thresholds for the Endangered category (10–500 km 2) under criterion B2. (4) Geographic proximity (from 1.7 to 18.4 km between localities) and shared anthropogenic threats in all known localities merits considering all as a single threat-defined location. (5) There is ongoing habitat decline due to deforestation for mining operations, legal and illegal small and large-scale logging, and continued environmental pollution caused by mining projects. Currently available information suggests that the extinction risk of P. daquilemai may be relatively high and we propose that it should be classified under the IUCN threatened category Endangered (EN), based on criteria B1ab(iii) + 2ab(iii). The species have not been found in protected areas, but it may inhabit El Quimi, El Zarza, and El Condor biological reserves, in Cordillera del Condor. Other areas in the region are protected forests, but they are not national protected areas and receive little or no real protection (Prieto-Albuja et al. 2019). Research and monitoring actions should be established to study the ecology of P. daquilemai and other endemic species of the Cordillera del Condor.
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